www.thaiagj.org Thai Journal of Agricultural Science 2008, 41(1-2): 11-18

Diversity of Soil-Litter : Comparison of the Pselaphine (Coleoptera: Staphylinidae: ) and the Ground Ants (Hymenoptera: Formicidae)

W. Sakchoowong1,*, W. Jaitrong2, K. Ogata3, S. Nomura4 and J. Chanpaisaeng5

1Forest Entomology and Microbiology Group, National Parks Wildlife and Plant Conservation Department, Bangkok 10900, Thailand 2National Science Museum, Klong 5 Klong Luang, Pathumthani, 12120, Thailand 3Institute of Tropical Agriculture, Kyushu University, Fukuoka 812-8581, Japan 4Division of Entomology, Department of Zoology, National Museum of Nature and Science Shinjuku, Tokyo 169-0073, Japan 5Department of Entomology, Faculty of Agriculture, Kasetsart University Bangkok 10900, Thailand

*Corresponding author. Email: [email protected]

Abstract

Forest litter-inhabiting except ants are poorly understood because of their small sizes and cryptic habitats. However, they play an essential role in nutrient cycling and contribute valuable data to study of comparative biodiversity and conservation. Soil beetles such as Pselaphine beetles (Coleoptera: Staphylinidae: Pselaphinae) are cosmopolitan, species-rich, and yet poorly studied, particularly in the tropics. While ants (Hymenoptera: Formicidae) are a large component of the community on ground in forest ecosystems. We sampled pselaphine beetles and ants in three types of primary forest and two types of disturbed forest habitat in eastern Thailand to assess the utility of pselaphine beetles and ants to determine if one group could be predicted the diversity of other. We compared species richness, abundance and diversity of ground ants and pselaphine beetles in each forest types using 1 m2 quadrat samples and extracted them with Tullgren funnels. A total of 1,867 pselaphine individuals representing 114 species in six supertribes and 44,135 individuals, 142 species in nine subfamilies of ants were collected in all habitats. Species richness, abundance, and Shannon diversity of pselaphine beetles and ants showed a significant response to habitat types as the same manner. Both pselaphine and ant species, abundance and diversity were significantly lower with increasing habitat disturbance. Strong relationships pattern between the two groups were significantly correlated in species richness, number of individuals and Shannon’s diversity index (H’) except Pielou’s evenness index (J’). In this study, we found that the relationships of ants and pselaphine beetles could be predicted the change in species richness, abundance and diversity of each other in response to habitat types.

Keywords: diversity pattern, pselaphine beetles, ants, soil and litter, Eastern forest of Thailand

Introduction 1992; Schulze et al., 2004). However, litter-inhabiting insects are poorly studied groups because of their A number of studies on biodiversity have small size and complex habitats. Although, they been increasing in the tropical rainforests (Wilson, play an essential role in soil i.g. nutrient cycling, 12 W. Sakchoowong et al. Thai Journal of Agricultural Science bioturbation and contribute valuable data to studies 58.31 km2. Precipitation in KARN is blocked by of comparative biodiversity and conservation high mountain of KSD which acts as a natural (Coddington et al. 1997; Stork, 1988). Pselaphine fence, making the climate relatively drier than those beetles (Coleoptera: Staphylinidae: Pselaphinae) in KSD and KKK. The annual precipitation in are among the most species-rich forest litter- KARN and KSD are relatively different with 1,600 inhabiting group (Carlton, 1999) and they play an mm in KARN and 3,800 mm in KSD. In eastern important role in the soil ecosystem as predators of Thailand, dry season lasts from November to small invertebrates (Newton and Chandler, 1989). March and wet season lasts from May to October This subfamily is species-rich and cosmopolitan. (Thailand Meteorological Department, 2007). Pselaphine beetles might therefore be profitably Differences of local precipitation and the altitude of used as indicator species for identifying habitat eastern forest areas ranging from 100 to 1,675 m differences and lack of disturbance in old-growth produce three various forest types (Kasetsart forests (Newton and Chandler, 1989). University, 2007), mixed deciduous forest, moist While, the ground ants perform a wide range of evergreen forest and hill evergreen forest. In ecological function as predators, preys, or addition for considering of human interruption mutualists with plants or (Schultz and which might be effect to local diversity of McGlynn, 2000). They are relatively sensitive and pselaphine beetles, we chose two inferior forest responsive to changes of ecosystem in habitats both types: secondary mixed deciduous forest (SMDF) spatial and time and they have been using as indicator and teak plantation forest (TP) for our experiment. taxon to assess impacts of environmental change on The detailed descriptions of forest types are shown biodiversity (Agosti et al., 2000; HÖlldobler and as following: Wilson, 1990; Toda and Kitching, 1999). In general, any forest conversions to other land Primary mixed deciduous forest (PMDF) use types will have markedly negative impacts on PMDF was dominated by Lagerstroemia venusta, diversity of animals and plants. Most studies just Lagerstroemia cuspidata, Pterocarpus macrocarpus, quantified the impact on diversity of single group Suregada multiflorum and Syzygium pergamenta- (Chey et al., 1997; Stork et al., 2003; Watt et al., ceum. Mixed deciduous forest site was established 2002). Only few studies had been reported the at Lum Jang Wat sub-district in KARN (N 13° 16' impact of tropical forest modification on several 25'', E 101° 44' 44'', 155 m elevation). taxonomic groups (Lawton et al., 1998; Schulze et al., 2004), however, their studies showed that no Moist evergreen forest (MEF) strongly pronounced relationships of change in MEF site was established at Ban Thung Krang species richness among the different groups. sub-district (N 13° 01' 08'', E 102° 12' 46'', 329 m Because of the importance of our understanding elevation). The dominant tree species are Diptero- of diversity pattern relationship of insect groups, carpus alatus, Mallotus peltatus, Shorea guiso, Strom- we undertook studies on species richness and bosia javanica and Diospyros transitoria. abundance of the pselaphine beetles and the ground ants in natural habitat. The results are reported in Hill ever green forest (HEF) this paper. HEF site was established at Khao Prabad (N 12° 50' 14'', E 102° 10' 20'', 1,069 m elevation). The Materials and Methods most abundant tree species in HEF were Scaphium scaphigerum, Castanopsis piriformis, Archidendron Study Sites quocense and Hornstedtia glabra. The eastern forest complex of Thailand consists of two wildlife sanctuaries, Khao Ang Rue Nai Secondary mixed deciduous forest (SMDF) Wildlife Sanctuary (KARN) with total area 1,030 Thirty percent of the land area in the northern km2, Khao Soi Dao Wildlife Sanctuary (KSD) with part of KARN was disturbed by human settlements total area 744.58 km2 and one national park, Khao and agriculture in 1992. The Royal Forest Department Kitchakut National Park (KKK) with total area has since relocated the villages and restored these Vol. 41, No.1-2, 2008 Diversity of soil-litter insects 13 areas. The forest has been recovering for approximate- Sorting and Identification ly 15 years. In the time of sampling, most of the The samples were taken to the laboratory, areas once opened by agriculture and villages are pselaphine beetles and ants were separated and stored covered with vegetation, and some of these areas in 80% ethanol. Pselaphine specimens were identified are now approaching an advanced stage of to genus using the keys of S. Nomura (S. Nomura, secondary forest; trees are approximately 10-12 m unpublished identification guide on subfamilies apart with diameter of 8-15 cm. We sampled an Protopselaphinae and Pselaphinae of Asia [Staphyli- SMDF site at Phuthai sub-district (N 13° 24' 56'', E nidae: Protopselaphinae; Pselaphinae] 2006), which 101° 52' 53'', 101 m elevation). The most abundant adopts the taxonomic conventions of Newton and tree species were S. multiflorum, L. venusta, S. Thayer (1995) and Chandler (2001). After generic pergamentaceum and P. macrocarpus. identification, specimens were then identified to morpho-species based on the external appearance Teak plantation (Tectona grandis L.) (TP) of specimens–a technique commonly used as a TP (Tectona grandis L.) were established in the surrogate for species level identification in boundary on the western part of KSD from 1969 to biodiversity studies (Abbott et al., 2002). 1992 by the Soi Dao Seed Orchard Station under Ant specimens were identified to genus using the Royal Forest Department project (Soi Dao key of Bolton (1994) and Hashimoto and Rahman Forest Seed Orchard Station, 1992). The plantations (2003), and later to species and morpho-species level, covering 376 ha have been designated for research using the reference collection of Thailand National and economic plantation. All teak plantations are Science Museum (TNSM). Voucher specimens of connected forming one large monoculture habitat in pselaphine beetle were mounted, labeled, and the KSD region. The teak plantation site (N 12° 58' deposited in the Forest Insect Museum (FIM), 48'', E 102° 17' 49'', 202 m elevation) chosen for Department of National Parks, Wildlife and Plant this study was located within a 20 ha area planted Conservation (DNP), Bangkok, Thailand. Voucher in 1976 with spacing of 6x6 m. The average specimens of ant were mounted, labeled and diameter (DBH) of teak trees was 30 cm and the deposited in the Forest Insect Museum (FIM) and average height was 25 m. Other tree species in the Thailand National Science Museum (TNSM). study site were Parkia sumatrana Miq. subsp. streptocarpa (Hance), P. macrocarpus Kurz, Data Analysis Lepisanthes rubiginosa (Roxb.) and Dalbergia Parameters of pselaphine beetles and ants: nigrescens Kurz. number of species and individuals, Shannon’s diversity index (H’) and Pielou’s evenness index Sampling Method (J’) from six samplings during 2006 were analyzed Ten 1 m2 quadrats were randomly sampled in and contrasted. Species, individuals, Shannon’s each site. The litter and surface soil to a depth of 3 diversity index (H’) (Magurran, 1988), and Pielou’s cm was scraped from the ground and sifted through evenness index (J’) (Ludwig and Reynolds, 1988) a 1 cm wire mesh sieve to exclude larger debris. were calculated. The Kruskal-Wallis nonparametric After sifting, each sample was transferred to a test (H) was conducted to test differences in beetle debris bag for transportation to the field station and ant species, individuals, Shannon’s diversity where Tullgren funnels were set up. After arriving index (H’) and Pielou’s evenness index (J’) among at the field station, the contents of each in debris habitats. Relationship between the parameters on bags were weighed and divided into ten samples, number of species, individuals Shannon’s diversity then put in Tullgren funnels for extraction. Soil and index (H’) and Pielou’s evenness index (J’) of ants litter dwelling organisms were extracted over 48 h and pselaphine beetles were tested using Spearman’s using 60 Watt incandescent lights. Specimens were correlation coefficient (r) and regression. Systat preserved in 80% alcohol prior to processing. Version 8 program (Systat Software, 1998) was Collections were conducted at bimonthly intervals used for Kruskal-Wallis nonparametric test (H), during the months of January, March, May, July, Spearman’s correlation and regression analysis. September and November 2006. 14 W. Sakchoowong et al. Thai Journal of Agricultural Science

Results 50 45 Ants Pselaphines Diversity Pattern between Pselaphine Beetles 40 35 and Ground Ants 30

A total of 44,135 individuals, 142 species in 25 nine subfamilies of ants were collected in all 20 habitats in 2006. The total number of ant species in species of Number 15 2006 in each habitat from highest to lowest were as 10 following: MEF (85 species), followed by 77 species 5 0 in PMDF, 68 species in SMDF, 61 species in HEF MEF PMDF HEF SMDF TP and 56 species in TP. While the total number of Forest habitat pselaphine beetles had lower with 1,867 individuals and 114 species. However, pselaphine species rank Figure 1 Pselaphine beetles and ant species richness in five different forest habitats in eastern Thailand. Bars was slightly different from the rank number of ant show mean ± SE (n = 6). MEF, moist evergreen forest; species in habitats as SMDF had situated in the PMDF, primary mixed deciduous forest; SMDF, secondary fourth rank as followed; MEF (66 species), PMDF mixed deciduous forest; HEF, hill evergreen forest; TP, (42), HEF (42), SMDF (23) and TP (21). teak plantation. Mean number on species richness of ants and pselaphines showed a significant respond to habitat 4.0 Ants Pselaphines types (Kruskal-Wallis one-way analysis of variance 3.5 for pselaphine beetles, H = 18.51, P < 0.01; ants, H ) 3.0 og

= 21.17, P < 0.01). Pselaphine and ant species (l 2.5 ance ance

richness decreased from MEF to TP (Figure 1). d 2.0 un Both pselaphine beetle and ant abundances were b

ve a 1.5 ti

also significantly greater in primary forests (MEF, a l e PMDF and HEF) than those in the secondary forest R 1.0 and teak plantation (pselaphine beetles, H = 23.04, 0.5 P < 0.01; ants, H = 20.62, P < 0.01; Figure 2). 0.0 MEF PMDF HEF SMDF TP Abundance of pselaphine beetles and ants in Forest habitat primary forests particularly PMDF and MEF was several times higher than in disturbed forest Figure 2 Pselaphine beetles and ant abundances in five habitats (SMDF and TP). different forest habitats in eastern Thailand. Mean ± SE Shannon’s diversity (H’) indices of pselaphine (n = 6). See site abbreviations in Figure 1. beetles and ants were higher in primary forest habitats than those in disturbed habitats. Moist 3.5 evergreen forest had highest diversity and Ants Pselaphines 3.0 decreased from PMDF to TP (Figure 3). Significant difference in Shannon’s diversity indices were 2.5 found among habitats for both taxa (pselaphine 2.0 beetles, H = 13.35, P < 0.01; ants, H = 16.35, P < 1.5

0.01). indexShannon's 1.0 However, the Pielou’s evenness indices (J’) which expresses the equality of species in each 0.5 habitat was not significant difference for ants (H = 0.0 1.78, P = 0.77) but it was significant difference for MEF PMDF HEF SMDF TP Forest habitat pselaphine beetles (H = 10.21, P < 0.05) (Figure 4). Pielou’s evenness index (J’) for pselaphine beetles Figure 3 Shannon’s index (H’) of pselaphine beetles and was higher in SMDF and TP than those in primary ant in five different forest habitats in eastern Thailand. forests. Bars show mean ± SE (n = 6). See site abbreviations in Figure 1. Vol. 41, No.1-2, 2008 Diversity of soil-litter insects 15

Correlation between Pselaphine Beetles and Ground Ants

Correlations of ants and pselaphine beetles were 1.2 undertaken between the variables on species Ants Pselaphines richness, individuals, Shannon’s diversity index 1.0

(H’) and Pielou’s evenness index (J’) and also 0.8 separately analyzed for different season (Table 1). There was strong relationship of parameters 0.6

between the two taxa. Almost parameters between evenness Pielou's 0.4 pselaphine beetles and ants were significantly 0.2 correlated except Pielou’s evenness index (J’) which had not significantly correlated (Figure 8, 0.0 Table 1). All these relationships had a Spearman’s MEF PMDF HEF SMDF TP Forest habitat correlation coefficient (r) higher than 0.51 (51%) and there were correlated both parameters in pooled Figure 4 Pielou’s evenness index (J’) of pselaphine data and separated data sets (Table 1), i.e. species beetles and ant in five different forest habitats in eastern richness between pselaphine beetles and ants in Thailand. Bars show mean ± SE (n = 6). See site pooled data were correlated at 62% (r = 0.619, P < abbreviations in Figure 1.

0.001, Figure 5), abundance (77%) (r = 0.768, P < 0.001, Figure 6) and Shannon diversity index (H’) 45 (51%) (r = 0.515, P < 0.001, Figure 7) and reached y = 0.7553x - 11.703 a level of highly significant (P < 0.01) and can be 40 35 described sufficiently by a linear regression model 30

(Figures 5-7). 25

20 Discussion 15 10 Number of species (pselaphines) of species Number Species richness, abundance, and Shannon’s 5 0 diversity of pselaphine beetles and ants showed a 15 20 25 30 35 40 45 50 55 Number of species (ants) significant response to habitat types. Both pselaphine beetle and ant diversity were significantly lower with increasing habitat disturbance as followed, MEF > Figure 5 Relationship between number of species of pselaphine beetles and ants across the habitat gradient (n PMDF > HEF > SMDF > TP. = 30). The patterns of species richness and diversity of these two taxa had dramatically responded to habitat types as the data showed that primary forests 400 y = 0.0604x - 26.91 (MEF, PMDF and HEF) hold the higher species 350 numbers and Shannon’s diversity index (H’) than in 300 SMDF and TP. Almost parameters except Pielou’s 250 evenness index (J’) showed strongly correlation in 200 pooled data set (all season) and also found in 150 separate data sets between dry and wet season 100 50

(Table 1). Number of individuals (pselaphines) 0 Interestingly, there was habitat specific for 0 1000 2000 3000 4000 5000 pselaphine beetles as that found in evenness value Number of individuals (ants) which had higher index (J’) in secondary forest and teak plantation than those in the primary forests, but Figure 6 Relationship between number of individuals of in ants there were not significantly different in pselaphine beetles and ants across the habitat gradient (n evenness index among primary forests and degraded = 30).

16 W. Sakchoowong et al. Thai Journal of Agricultural Science

Table 1 Relationship (Spearman’s correlation) between pselaphine beetles and ants surveyed across sites, the top and right portion of the table presents correlations of parameters (Spearman’s (r)) and the number of samples available for pairwise comparison, \n (in square brackets).

Dry season Wet season All season (15) (15) (30)

Species richness 0.600** 0.819*** 0.619***

Individuals 0.879*** 0.627** 0.768***

Shannon's diversity (H’) 0.581* 0.510* 0.515**

Pielou's evenness (J’) 0.205 -0.168 -0.70 Level of significance * = P < 0.05, ** = P < 0.01, *** = P < 0.001

forests. This is indicated that pselpahine beetles are 3.0 y = 1.0752x - 1.1505 more spatially homogenous in degraded forests while ants seem to be more ubiquitous (Andersen, 2000). 2.5 Published papers on biodiversity indicator of

2.0 different groups covering a diverse spectrum of taxonomic groups across a similar land use gradient 1.5 also reported a general trend of decreasing species richness with increasing habitat modification (Lawton

Shannon's index (pselaphines) 1.0 et al., 1998; Schulze et al., 2004). However, there

0.5 were no strongly pronounced relationships of the 1.5 1.7 1.9 2.1 2.3 2.5 2.7 2.9 3.1 3.3 3.5 change in species richness among the different Shannon's index (ants) animal group surveys. This could be resulted from

Figure 7 Relationship between Shannon’s index (H’) of different taxa would response to the disturbance in pselaphine beetles and ants across the habitat gradient (n different ways. Different groups such as birds, = 30). butterflies, dung beetles, ants, spiders, etc, have different guilds and live in spacious habitat, thus they

would response to different degrees to habitat alteration 1.10 and all these animals were sampled in different

1.00 methods (Lawton et al., 1998; Schulze et al., 2004). On the other hand, in this study, there were 0.90 positively significant correlated of most parameters 0.80 between ants and pselaphine beetles (except Pielou’s 0.70 evenness index) and the Spearman’s correlation 0.60 coefficient (r) among parameters were high. These Pielou's index(pselaphines)

0.50 strong relationships between ants and pselaphine beetles in this study might be reasoned as (1) both 0.40 0.60 0.65 0.70 0.75 0.80 0.85 0.90 taxa have similar natural niches (factors necessary for Pielou's index (ants) species existence) (Molles, 2005) i.e. they have similar resource use and similar habit (guild), and they live in Figure 8 Relationship between Pielou's evenness (J’) of pselaphine beetles and ants across the habitat gradient (n similar microhabitat (soil and litter). (2) and many = 30). pselaphine genera in several supertribes plus supertribe

Vol. 41, No.1-2, 2008 Diversity of soil-litter insects 17

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